Experimental alterations of the spores of lycopodium clavatum as related to diagenesis

Lycopodium clavatum spores have been heated to different temperatures at atmospheric pressure, at room temperature with 0.5 kbar pressure and at different temperatures with 1 kbar pressure, The effects of heat, pressure, and heat and pressure together on the spore have been examined in detail using different microscope techniques. Effects of some chemicals on these spores have also been observed.It is known that temperature and pressure change the colour of spore and pollen grain walls, mainly the exine (outer wall of the spore). Changes to the intine and the matter present in the cytoplasmic cavity (= inner contents), however, have not been taken into account by very many workers. In most of the previous works the inner contents were extracted before the experimental treatment began.In the present work, unextracted spores are used for the experiments which show two types of alterations of the spores with rising temperature at atmospheric pressure: (1) alteration of the inner contents, i.e. gradual colour change of the inner contents and their ultimate exudation from the spore through the exine at about 300°C; (2) gradual shrinkage of the exine due to the exudation of the inner contents which also causes an overall size reduction of the spore from 200° to 350°C. The exine does not change its colour up to 325°C; and this temperature, its starts to change its colour slowly, amalgamates with other exines of the empty spores, becomes amorphous, and ultimately deteriorates into unidentifiable organic matter.The process of colour change of the inner contents of spores and the general deformation are much slower when the spores are subjected to 1 kbar pressure with rising temperatures. Spores at room temperature with 0.5 kbar pressure show no colour change but only physical deformation.     

Re-Exposure of Tapes at High Temperature and Pressure in the Lycopodium Clavatum Spore Exine

Lamellations are visualizable through the staining commonly used in transmission electron microscopy during exine formation on Lycopodium and other spores, and the nexine of pollen grains. The lamellations so exposed consist-of dark tapes at either side of an unstained (white) line. Neither tapes nor white lines are visualizable in the exine of mature spores of Lycopodium. The continued presence of lamellations having tape-white line spacing has been demonstrated with inorganic tracers in the nexine of pollen in which lamellations otherwise appeared to be absent. Through high contrast staining methods for TEM we have observed lamellations in the residual exine following heat treatment (350[ddot]C) of mature spores of Lycopodium clavatum. The surface of these residual exines was etched by treatment with hot 2-aminoethanol and filaments were observed to protrude from the etched surfaces. The residual exine stained darkly. Lycopodium spores heated to 300[ddot]C at 1 kb pressure had long filaments exposed at the surface of the residual exine (sporopollenin). Sections of the pellet remaining after heat and pressure treatment also included bundles of closely parallel filaments and masses of isolated filaments. The filaments were stained while the exine residue, assumed to include sporopollenin, was not. Isolated filaments produce a stable metachromasia with toluidine blue indicating the presence of many closely spaced sites of negative charge. The staining of intact exines with basic dyes may result from anionic sites on filaments embedded within the exine rather than being due to sporopollenin. The results of our experiments indicate that the filaments are more resistant to heat and pressure and 2-aminoethanol than is sporopollenin. We propose that the trilamellar elements commonly called tapes and white lines might be composed of two filaments bridged by polybasic molecules.     

STUDIES OF SEED FERN POLLEN: DEVELOPMENT OF THE EXINE IN MONOLETES (MEDULLOSALES)

Monoletes pollen extracted from the seed fern synangium Dolerotheca sclerotica Baxter illustrate four stages in the development of the sporoderm. In the first stage the grains are up to 100 μm long and possess an apparent homogeneous exine in which there is little differentiation between the nexine and sexine. Numerous nexine lamellae and the initiation of sexine expansion mark stage 2 in exine ontogeny. Further expansion of the sexine continues in the third stage until the ratio between the nexine and sexine is approximately 1:5. The final stage in maturation of the sporoderm shows an expanded alveolate sexine with some of the sporopollenin units broken and disorganized. It is at this stage of development that nexine lamellae are most prominent. The formation of sporoderm layers in the fossil grains is compared with pollen grain development in living cycads (Cycadophyta) and a model proposed to account for the apparent early formation of nexine lamellae in Monoletes. The evolution of exine components in early pollen types is discussed.     

中国芸香科植物的花粉形态

本文用光学显微镜对我国21属52种2变种芸香科(Rutaceae)植物的花粉形态进行了详细地观察研究,其中对6属6种进行了扫描电镜观察。本科花粉具沟孔,是花粉单类型的科之一。根据萌发孔的数目和外壁纹饰特点,本文将芸香科的花粉分为具3孔沟和4-6孔沟两个花粉类型,并根据各属花粉形态特征,作出了花粉形态分属检索表。此外,本文还对花椒(Zanthoxylum L.)和Fagara两属的合并问题及花椒与吴茱萸(Evodia Scop.)两属之间的亲缘关系等问题进行了讨论。     

SOLUBILITY OF POLLEN EXINES

The sporopollenin of pollen exines of Ambrosia trifida is soluble in fused potassium hydroxide, in strong oxidizing solutions, and in certain organic bases. It is insoluble in other organic and inorganic acids and bases, in lipid solvents, and in detergents. The outer exine layer of gymnosperm and angiosperm pollen dissolves in 2-aminoethanol. The inner exine layer, as well as the exine of pteridophyte spores, is insoluble. The exine dissolution process in 2-aminoethanol involves swelling and disintegration of exine structures, leaving some residual globules. Sporopollenin shares some solubility properties with lignin and cutin but appears to be chemically distinct from these substances.     

THE ULTRASTRUCTURE OF IN SITU CLAVATIPOLLENITES POLLEN FROM THE EARLY CRETACEOUS OF PATAGONIA

Small dispersed anther contents of Clavalipollenites grains are reported from the Early Cretaceous (early Aptian) of Patagonia. This report represents the first documentation of in situ grains of this type from the Southern Hemisphere. The anthers are small (0.6 mm long × 0.25 mm wide); no tissues of the androecium are preserved nor is there any indication of how the sacs were attached. Some grains are still in tetrads and closely associated with numerous orbicules and tapetal membranes. Grains are 18–22 μm long and up to 15 μm wide. The exine consists of an inner homogeneous nexine that supports narrow columellae below a perforate tectum. The mature pollen wall includes uniform microgranules that ornament the muri. The chloranthaceous affinities of these Gondwana pollen sacs are established and the grains are compared with specimens recovered from slightly older or coeval sediments from the Northern Hemisphere. The discovery of these pollen sacs from Patagonia expands our understanding of early angiosperm biogeography.     

Pollen Morphology of Loxostemon (Cruciferae) in China

The present paper deals with the pollen morphology of 10 species and 1 variety of Loxostemon in China. The pollen grains were all examined under light microscope. The pollen grains of Loxostemon are subspheroidal, spheroidal or prolate, 18--33×11.8-28 μ in size, 3-colpate, colpi 15-21 μ long and 1-2 μ wide. The exine is 1.5-3 μ thick with two indistinct or distinct layers. All the pollen grains are generally reticulate under light microscope. They are distinctly or obscurely and finely reticulate. L. axillus and L. repens are generally similar in gross morphology, but the pollen grains of these two species are different. The pollen grains of L. axillus are regularly polygonally reticulate, colpi are acute-ended and the exine is about 3 μ thick, whereas those of L.repens are irregularly polygonally reticulate, colpi are enlarged at both ends and the exine is about 2.8 μ thick. L. incanus and L. stenolobus appear to have similar gross morphology, but the pollen grains of the former have exines with two distinct layers and a densely and finely reticulate ornamentation and those of the latter have exines with two indistinct layers and a flexuosely reticulate ornamentation.     

Distyly in Polygonum jucundum Meisn. (Polygonaceae)

Pollen grains of tribe Sanguisorbeae (Rosaceae, Rosoideae) were examined using scanning electron microscopy to identify useful characters, test taxonomic and phylogenetic hypotheses among genera, and elucidate pollen character evolution based on a molecular phylogeny. Aperture number, aperture structure, pollen shape, and exine sculpturing were variable within Sanguisorbeae and were used to delineate six pollen types. Four types (I–IV) were observed only in subtribe Sanguisorbinae whereas two types (V–VI) were found only in subtribe Agrimoniinae. Pollen grains of tribe Sanguisorbeae were generally subprolate to spheroidal in shape, had operculate or pontoperculate apertures, and had three apertures, except for Margyricarpus (tetraperturate). Exine sculpturing within Sanguisorbinae represented variations of striate, verrucate, rugulate, and perforate patterns often with microechinate sculpturing. Striate exine patterns and prolate shapes characterized the pollen of the Agrimoniinae, except for the microechinate-verrucate pattern and subprolate to spheroidal shapes observed in Hagenia. Pollen characters are most useful at the generic level and, when mapped on to a molecular phylogenetic tree of the tribe, are concordant with a monophyletic Agrimoniinae and a clade comprising Margyricarpus + Acaena + Polylepis + Cliffortia + Sanguisorba in the Sanguisorbinae. Outgroup comparison indicated that operculate colpi, three apertures, and polymorphism for striate or microverrucate exines represented primitive states for tribe Sanguisorbeae.     

Seed set,pollen morphology and pollen surface composition response to heat stress in field pea

Pea (Pisum sativum L.) is a major legume crop grown in a semi‐arid climate in Western Canada, where heat stress affects pollination, seed set and yield. Seed set and pod growth characteristics, along with in vitro percentage pollen germination, pollen tube growth and pollen surface composition, were measured in two pea cultivars (CDC Golden and CDC Sage) subjected to five maximum temperature regimes ranging from 24 to 36 °C. Heat stress reduced percentage pollen germination, pollen tube length, pod length, seed number per pod, and the seed–ovule ratio. Percentage pollen germination of CDC Sage was greater than CDC Golden at 36 °C. No visible morphological differences in pollen grains or the pollen surface were observed between the heat and control‐treated pea. However, pollen wall (intine) thickness increased due to heat stress. Mid‐infrared attenuated total reflectance (MIR‐ATR) spectra revealed that the chemical composition (lipid, proteins and carbohydrates) of each cultivar's pollen grains responded differently to heat stress. The lipid region of the pollen coat and exine of CDC Sage was more stable compared with CDC Golden at 36 °C. Secondary derivatives of ATR spectra indicated the presence of two lipid types, with different amounts present in pollen grains from each cultivar.     

Dimorphic exine sculpturing in two distylous species ofDyerophytum (Plumbaginaceae)

Light and SEM observations on the pollen ofDyerophytum africanum andD. indicum have revealed marked differences in exine features. These distylous species also have dimorphic pollen. In the short-styled individuals of both species, the sexine and nexine are of equal thickness, and the clava-like sexinous processes are short without marked projections. In the long-styled individuals, the sexine is thicker than the nexine, the clavae are higher than broad with an apical spinule. Pollen size and apertures are identical in both morphs. — Palynological evidence is presented for relationships betweenDyerophytum andCeratostigma, Plumbago andAegialitis. Moreover, the genusDyerophytum exhibits pollen morphological similarities with some species ofLinum (Linaceae).     

Pollen ultrastructure in <Emphasis Type="Italic">Aristolochia manshuriensis</Emphasis> and <Emphasis Type="Italic">A. contorta</Emphasis> (Aristolochiaceae)

Pollen ultrastructure has been studied in two relict and rare species of the genus Aristolochia, A. contorta Bunge and A. manshuriensis Kom. (Aristolochiaceae). Both species have inaperturate, spheroidal, sometimes distally monocolpate or distally bicolpate pollen grains. The equatorial and polar axes of pollen grain in A. manshuriensis are 48.5 and 44.0 μm, respectively. The percentage of defective pollen grains in A. manshuriensis is 3.4%. The fossulate, perforated exine is up to 2.3 μm in thickness; the sexine and the nexine are almost equal in thickness. In A. contorta, the equatorial axis of pollen grain is 36.6 μm: the defectiveness percentage, 24.5%. The exine is verrucate, up to 0.3 μm in thickness, while the sexine is two to three times thicker than the nexine. The pollen germination experiments have shown that pollen of A. manshuriensis, in contrast to A. contorta, can germinate in 10–20% sucrose at 22°С. These data and the high percentage of pollen defectiveness in A. contorta indicate that the androecium function in this species is reduced. The reduction of the androecium function is evidenced by a small amount of pollen grains in anthers or empty anthers and a high percentage of defective pollen grains.     

A new elater-bearing Late Albian pollen species from offshore eastern Saudi Arabia

A new elater-bearing pollen referable to Elaterocolpites is described from Late Albian sediments of the Arabian Gulf, offshore eastern Saudi Arabia. This pollen has a main body consisting of five elater-like projections in the equatorial area and five colpi, one colpus at the base of each projection. The exine has three layers. The foot-layer forms the nexine whereas the other two layers, the baculate-layer and the tectum, form the sexine. The exine structure has an angiosperm affinity. The pollen occurs within the Middle Cretaceous phytogeoprovince characterized by its elater-bearing palynomorph occurrences.     

Study on the Pollen Morphology of the Chinese Family Oleaceae

This paper deals with the pollen grains of 11 genera, 22 species in Chinese Oleaceae. A comparative study on the pollen grains was carried out by means of scanning electron microscope and light microscope. The pollen grains of the family Oleaceae are almost spheroidal, oblate and prolate, in polar view tlie majority of the pollen grains are tri-lobulate-circular, the minority are tetra-lobulate-circular and dilobulate-circular. Pollen size: (15-65) × (12- 60)μ, 3-colporate and 3-colpate the minority of the pollen grains 4-colpate and 2- colpate colpus usually distinct and the en doaperature vague. The exine oonsists of two layers, the sexine thicker than the nexine, with big reticulum or fine reticulum, the majority of the reticulum are very distinct. On the basis of the pollen size and exine ornamentation, the pollen grains are divided into following two main types: (1) The larger pellen grains and the exine with big reticulum are found in the genera: Jasminum, Nyctanthes, Ligustrum, and Syringa. (2) The smaller pollen grains and the exine with finer reticulum are found in the genera: Osmanthus, Olea, Chionanthus, Fraxinus, Forsythia, Fontanesia and Linociera. In view of the features of pollen morphology, this family is much similar to the family Itamamelidaeeae. For example, the first pollen type resembles the genus Coryiopsis of the family Hamamelidaceae, but there are some differences between them, the former with a bigger reticulate exine and without any colpal membrane, the latter with a finer reticulate apocolpium and eolpate membrane. The second pollen type is much similar to that of the genera Hamamelis and Loropetalum of family Hamamelidaceae.     

Pollen morphology of the subfamily Aurantioideae (Rutaceae)

The Aurantioideae is one of seven subfamilies of the Rutaceae consisting of two tribes, the Clauseneae, containing five genera, and the Citreae, with 28 genera. Each tribe contains three subtribes. The pollen morphology of the subfamily Aurantioideae is described and illustrated for the first time based on light and scanning electron microscopy. Five pollen types have been recognised in the subfamily, based mainly on aperture number and exine ornamentation. The pollen grains show a high degree of intergeneric variation. Pollen grains of Clauseneae are 3-colporate, microstriate or microstriato-reticulate, whereas pollen grains of Citreae are almost always 4/5 colporate with exines varying from microperforate to coarsely reticulate. Congruence between pollen types and the currently accepted classification is discussed, as well as the systematic implications of pollen morphology for the subfamily.     

Zur Pollenmorphologie der Gattung Waltheria L. (Sterculiaceae)

The pollen grains of 69 specimens representing 44 species or supposed species of the genus Waltheria L. have been studied. Based mainly on differences in exine characters 2 pollen types, a spinulose and a suprareticulate one were pointed out. Both types show ± parallel trends in the development of the aperture system to an increase of aperture number, to a shortening of the colpi and to more distinct bounds and nexine thickenings of the ora. They exemplify important steps of aperture evolution, which in the spinulose type leads from zonocolporate by way of zonopororate to pantopororate and in the suprareticulate type from zonocolporate to pantocolporate grains. The tendencies of sporoderm evolution concern a more distinct morphological and partially also a regional differentiation of the exine elements (bacula, spinulae), which prove to be close correlated with the evolution levels of the apertures. According to the pollen morphological point of view the species of the genus Waltheria are to be arranged in two groups, which may be regarded as two related, but relative independent phyletic entities. These evidences speak in favour of a taxonomic     

Pollen morphology of the genus Euphrasia L. (Orobanchaceae) from Pakistan and Kashmir and its taxonomic significance

Pollen morphology of 25 species of the genus Euphrasia L., belonging to the family Orobanchaceae from Pakistan has been examined by light and scanning electron microscope. It is a stenopalynous genus. Pollen grains are usually radially symmetrical, isopolar, tricolpate, sub-prolate to prolate-spheroidal rarely oblate-spheroidal. Sexine slightly thinner than nexine or as thick as nexine. Tectum coarse to fine retipilate or rugulate–retipilate. On the basis of exine ornamentation three distinct pollen types viz., Euphrasia foliosa, Euphrasia multiflora, and Euphrasia incisa are recognized. Pollen morphology is significantly helpful at specific level within the pollen type.     

中国壳斗科栎亚科花粉形态研究

本文应用光学显微镜与扫描电子显微镜观察了壳斗科所属栎亚科2属26种2变种的花粉形态,并以本科其它属一些代表种作对比,试图从孢粉学方面为栎亚科的系统分类提供依据。 花粉外壁纹饰在扫描电镜下可分三种类型,即水青冈型(含水青冈属),栎型(含栎属和三棱栎属),栗型(含栗属、锥属和柯属),花粉的类型与植物形态分类的三个亚科一致。 栎亚科的花粉外壁纹饰在扫描电镜下为颗粒聚集成的种种形状。栎属可区分为四种类型,即颗粒状、颗粒——蠕虫状、聚颗粒及芽孢状,其中类型一、二、三为常绿种类,类型四为落叶种类。花粉纹饰在常绿与落叶种类之间有较明显的差异,而青冈亚属和栎亚属之间却没有界线。因此,栎属仍以包含两个亚属为宜。三棱栎属花粉纹饰则介于栎属的类型二与三之间。     

Correlation between pollen morphology and pollination mechanisms in the Hydrocharitaceae

The pollen morphology of 11 genera and 11 species of the Hydrocharitaceae and one species of the Najadaceae was studied using scanning and transmission electron microscopies, and the exine structures and sculptures are discussed in relation to pollination mechanisms and the molecular phylogeny. The pollen grains of the Hydrocharitaceae are spherical, inaperturate, and form monads or tetrads, while those of the Najadaceae are elliptical, inaperturate, and form monads. The entomophilous genera Egeria, Blyxa, Ottelia, Stratiotes, and Hydrocharis share pollen grains that have projections like spines or bacula. The anemophilous genus Limnobium has reticulate pollen grains. The hypohydrophilous genera Thalassia and Najas are characterized by pollen grains with reduced exine structures. The pollen-epihydrophilous genera Elodea and Hydrilla have tightly arranged small spinous pollen grains, and the male flower-epihydrophilous genera Enhalus and Vallisneria have reduced reticulate or gemmate exines. Character state reconstruction of the exine structures and sculptures using a molecular phylogenetic tree suggests that variation in the exine is generally correlated with the pollination mechanism; the selective pressures acting on the pollination mechanisms have reduced the exine structure in hypohydrophilous plants and resulted in various exine sculptures that are adapted to the different pollination mechanisms in entomophilous, anemophilous, and pollen-epihydrophilous plants.     

西瓜花粉壁超微结构与花粉ATP酶细胞化学定位

研究了西瓜花粉壁超微结构以及单核花粉液泡化时期ATP酶活性超微细胞化学定位。花粉壁的外壁分为外层和内层 ,外层包括覆盖层、基粒棒和基足层等三层 ,内层只包含一层。外层电子密度相对较小 ,内层电子密度相对较大 ;外层与内层之间有缝隙。ATP酶活性反应产物主要分布在细胞质基质、质体、内质网和花粉内壁中     

西瓜花粉壁超微结构与花粉ATP酶细胞化学定位

研究了西瓜花粉壁超微结构以及单核花粉液泡化时期ATP酶活性超微细胞化学定位。花粉壁的外壁分为外层和内层,外层包括覆盖层、基粒棒和基足层等三层,内层只包含一层。外层电子密度相对较小,内层电子密度相对较大;外层与内层之间有缝隙。ATP酶活性反应产物主要分布在细胞质基质、质体、内质网和花粉内壁中。